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研究生:陳盈豪
研究生(外文):Ying-Hao Chen
論文名稱:台灣白花蝴蝶蘭斑葉突變株形成之分子層次探討
論文名稱(外文):Molecular studies of variegated mutation in Phalaenopsis aphrodite subsp. formosana (Orchidaceae)
指導教授:廖培鈞蔡奇助蔡奇助引用關係
指導教授(外文):Pei-Chun LiaoChi-Chu Tsai
學位類別:碩士
校院名稱:國立屏東科技大學
系所名稱:生物科技系
學門:生命科學學門
學類:生物科技學類
論文種類:學術論文
論文出版年:2013
畢業學年度:101
語文別:中文
論文頁數:91
中文關鍵詞:台灣白花蝴蝶蘭斑葉差異性剪輯
外文關鍵詞:Phalaenopsis aphrodite subsp. formosanavariegated leafalternative splicing
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台灣白花蝴蝶蘭 (Phalaenopsis aphrodite subsp. formosana) 為蘭花育種重要的親本,目前有發現斑葉突變株 (variegated mautant),其葉片會產生綠、黃相間的嵌紋。本研究目的在探討造成台灣白花蝴蝶蘭綠、黃嵌紋的形成機制。透過雷射掃描共軛焦顯微鏡與穿透式電子顯微鏡觀察綠、黃組織間的葉綠素累積和葉綠體結構,發現黃色組織中的質體 (plastid) 累積大量油滴小粒 (plastoglobulum) 並無類囊體結構。利用蛋白質二維電泳 (two-dimensional gel electrophoresis, 2-DE) 分析植株葉片綠、黃色組織總蛋白質表現差異,以 LC/MS/MS 鑑定其身分,經蛋白質資料庫比對後,除了部分蛋白無法鑑定外,可以鑑定且與綠、黃組織間差異有關的蛋白質有東亞蘭嵌紋病毒 (Cymbidium mosaic virus, CymMV) 的病毒外鞘蛋白 (coat protein)、葉綠體熱休克蛋白 70-1 (heat shock protein 70-1, HSP 70-1) 以及光系統II 中釋氧複合體 (oxygen-evolving complex, OEC) 中的 PsbO 與 PsbP 次單元。以 RACE 技術釣取 PsbO 與 PsbP 基因全長發現,兩種基因的轉錄體皆具多型性且都有內隱子保留的變異型轉錄體,透過 real-time PCR 檢測正常轉錄型與變異型的表現量總和,發現綠、黃組織中 PsbO 與 PsbP 表現量皆相同;黃色組織中 CymMV 蛋白質外鞘表現量高於綠色組織 3 倍。經由 semi-quantitative RT-PCR 發現,PsbO 的正常轉錄體在黃色組織表現量較高;PsbP 的變異型轉錄體在黃色組織表現量較高。由此推測,黃色組織可能因累積較多病毒對 PsbO 與 PsbP 進行後轉錄基因調控,由於 PsbP 蛋白扮演穩定類囊體摺疊重要角色,造成黃色組織內因為無法形成正常的類囊體,導致葉綠素無法正常累積而形成斑葉。
Phalaenopsis aphrodite subsp. formosana, one of important parent on Phalaenopsis breeding.There is a variegated mutant of the species can be found. In this study, we will examine to reveal the mechanism of gene regulation for the variegated mutant. The cross section of the variegated leaf was conducted by confocal laser-scanning microscopy and transmission electron microscope. The cells of green sectors contain normal chloroplasts, cells in the yellow sectors contain plastids that lack pigments and thylakoid. In proteomic analysis, four differential expression proteins between the green and yellow sectors of leaves can be identified as CPHSP 70-1, coat protein of CymMV, PsbP and PsbO by two-dimensional electrophoresis and LC/MS/MS. The semi-quantitative reverse transcriptase PCR and real-time PCR were used to amplify cDNA products reverse transcribed from mRNA and compare express between PsbO and PsbP and full-length cDNA of the genes mentioned above were cloned by the method of RACE technology. A full-length cDNA of PsbO and PsbP were cloned by RACE analysis and we fund that both PsbO and PsbP have the variant transcripts caused by intron retention. The total expression of both functional and variant transcripts from PsbO and PsbP between green and yellow sectors showed no difference by real-time PCR. Moreover, the expression of CymMV coat protein in yellow sector is higher than green sector for three times. Based on semi-quantitative RT-PCR, we found that the expression of both functional transcription of PsbO and variant transcription of PsbP are higher in yellow sector. It is suggested that yellow sector might accumulate more virus which result in the post-transcriptional regulation on PsbO and PsbP. Because PsbP play an important role in the stability of thylakoid folding, we suggested that the negative regulation of PsbP will block the development of thylakoid in yellow sector. It causes the chlorophyll can not accumulate in the yellow sector, as a result, the variegated leaves are shown.
目錄
中文摘要 ............................................................................................................ I
Abstract.. .......................................................................................................... III
誌謝..................................................................................................................V
目錄..................................................................................................................VI
圖表目錄 ......................................................................................................... IX
1. 前言..................................................................................................................1
2. 前人研究....................................................................................................... 2
2.1台灣白花蝴蝶蘭 ........................................................................................ 2
2.2植物斑葉現象 ........................................................................................... 2
2.3質體的生合成與分化 ............................................................................... 5
2.3.1 原生質體 ( proplastids) ..................................................................... 5
2.3.2 葉綠體結構和型態 ............................................................................ 6
2.3.3 澱粉體的結構和形態 ........................................................................ 7
2.3.4 有色體的結構和形態 ........................................................................ 7
2.3.5 植物光合作用系統 ............................................................................ 8
2.4 葉綠素的生合成 ...................................................................................... 9
2.4.1 葉綠素結構之合成 .......................................................................... 10
2.5 植物基因調控 ........................................................................................ 11
3. 材料與方法 ................................................................................................. 16
3.1 實驗材料................................................................................................ 16
3.2綠色與黃色組織的切片觀察 ................................................................. 16
3.2.1 徒手切片 (free hand section) .......................................................... 16
3.2.2 光學顯微鏡觀察綠、黃組織 .......................................................... 16
3.2.3 雷射掃描共軛焦顯微鏡 (confocal laser scanning microscope, CLSM) 觀察綠、黃組織 ............................................................... 16
3.2.4 穿透式電子顯微鏡 (transmission electron microscope, TEM) 觀察綠、黃組織 ............................................................................. 16
VII
3.3 蛋白質體學分析 .................................................................................... 17
3.3.1二維電泳之蛋白質定量方法 ........................................................... 17
3.3.2蛋白質二維電泳 ............................................................................... 18
3.3.3膠體內蛋白酶水解 ........................................................................... 18
3.4 總RNA的萃取 (total RNA extraction) ................................................ 19
3.5 反轉錄酶反應 (reverse transcriptional polymerase chain reaction) ....... 20
3.6 聚合酶連鎖反應 (polymerase chain reaction) ...................................... 20
3.7 RACE (rapid amplification of cDNA ends) ............................................. 21
3.7.1 製備第一股 cDNA.......................................................................... 21
3.7.2 3’ 和 5’端 cDNA 的增殖反應 ................................................. 21
3.7.3 由瓊脂凝膠中回收cDNA片段 ...................................................... 22
3.8 接合作用 (ligation reaction) .................................................................. 22
3.9 轉型作用 (transformation) .................................................................... 23
3.10 藍白篩選與養菌 .................................................................................. 23
3.11 微量抽取質體 (plasmid) ..................................................................... 23
3.12核酸定序分析 ....................................................................................... 24
3.13 real-time PCR分析基因表現量............................................................ 24
3.13.1 合成單股cDNA ............................................................................ 24
3.13.2 專一性引子之設計 ........................................................................ 24
3.14 蝴蝶蘭 PsbP 與 PsbO 基因啟動子的選殖 ...................................... 24
3.14.1 植物DNA的萃取 ......................................................................... 24
3.15 Genome WalkerTM Kit的操作 .............................................................. 25
3.15.1 Genomic DNA行限制酶作用 ........................................................ 25
3.15.2 純化限制利用 PCR AdvancedTM PCR Clean Up System (VIOGENE, Taipei, Taiwan) 來純化 DNA ................................... 25
3.15.3 Genome WalkerTM Adaptor和切割過之 genomic DNA的接合 ... 26
3.15.4 以專一性引子PCR放大目標基因 ............................................... 26
4. 結果..............................................................................................................28
VIII
4.1 台灣白花蝴蝶蘭斑葉突變株之外部型態觀察 ..................................... 28
4.1.1 台灣白花蝴蝶蘭突變株之嵌紋現象之觀察 ................................... 28
4.1.2 穿透式電子顯微鏡觀察台灣白花蝴蝶蘭斑葉突變株綠色與黃色組織內葉綠體的結構 .................................................................. 28
4.1.3 顯微鏡觀察台灣白花蝴蝶蘭斑葉突變株綠色與黃色組織內部細胞色素分布 ................................................................................. 28
4.1.4 台灣白花蝴蝶蘭之斑葉突變株自交後型態觀察 ........................... 28
4.2 以蛋白質體學探討斑葉綠、黃組織間總蛋白質表現量差異 .............. 29
4.2.1 二維電泳分析綠色與黃色組織間蛋白質表現量差異.................... 29
4.2.2 二維電泳分析綠色與黃色組織間表現量差異蛋白質身分鑑定 .... 29
4.3 基因之選殖分析 .................................................................................... 29
4.3.1 退化性引子選殖 PsbP 與PsbO cDNA的中間片段 ..................... 29
4.3.2 PsbP 與 PsbO 基因全長之選殖 .................................................... 30
4.3.3 PsbP 基因上游序列基因啟動子之選殖 ......................................... 31
4.3.4 綠色及黃色葉面組織內之 PsbP、PsbO 與 CymMV 基因表現量比較 ......................................................................................... 31
4.3.5 綠色與黃色葉面組織內之 PsbO 不同轉錄體之差異性表現 ....... 32
4.3.6 綠色與黃色葉面組織內之 PsbP 不同轉錄體之差異性表現 ....... 32
5. 討論 ..................................................................................................................34
6. 結論 ..................................................................................................................41
參考文獻 ......................................................................................................... 66
作者簡介 ......................................................................................................... 91
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